Evidence supporting an intentional Neandertal burialat La Chapelle-aux-SaintsWilliam Rendua,b,1, Cédric Beauvalc, Isabelle Crevecoeurd, Priscilla Bayled, Antoine Balzeaue, Thierry Bismuthb,f,Laurence Bourguignong,h, Géraldine Delfourd, Jean-Philippe Faivred, François Lacrampe-Cuyaubèrec,Carlotta Tavorminac,i, Dominique Todiscoj, Alain Turqd,k, and Bruno Maureilled

aCenter for International Research in the Humanities and Social Sciences, Unité Mixte Internationale 3199, Centre National de la Recherche Scientifique(CNRS)–New York University, New York, NY 10003; bTravaux et Recherches Archéologiques sur les Cultures, les Espaces et les Sociétés, Unité Mixte deRecherche (UMR) 5608, CNRS, Université Toulouse Le Mirail, F-31058 Toulouse Cedex 9, France; cArchéosphère, F-33300 Bordeaux, France; dde la Préhistoire àl’Actuel: Culture, Environnement et Anthropologie, UMR 5199, Ministère de la Culture et de la Communication (MCC), CNRS, Université de Bordeaux, F-33400Talence, France; eDépartement de Préhistoire du Muséum National d’Histoire Naturelle, Équipe de Paléontologie Humaine, UMR 7194 du CNRS, F-75013Paris, France; fDirection régionale des Affaires culturelles du Limousin–Service Régional de l’Archéologie, MCC, F-87036 Limoges Cedex, France;gGrand Sud-Ouest, Institut National de Recherches Archéologiques Préventives, F-24260 Campagne, France; hArsScan/AnTet, UMR 7041, CNRS, MaisonArchéologie et Ethnologie (MAE), F-92023 Nanterre, France; iInstitut Català de Paleoecologia Humana i Evolució Social, Universitat Rovira i Virgili, E-43007Tarragona, Spain; jDépartement de Géographie, UMR 6266 Identité et Différenciation des Espaces, de l’Environnement et des Sociétés, Université de Rouen,F-76821 Mont Saint Aignan Cedex, France; and kMusée National de Préhistoire, F-24620 Les Eyzies-de-Tayac, France

Edited by Erik Trinkaus, Washington University, St. Louis, MO, and approved November 15, 2013 (received for review September 8, 2013)

The bouffia Bonneval at La Chapelle-aux-Saints is well known forthe discovery of the first secure Neandertal burial in the early 20thcentury. However, the intentionality of the burial remains an issueof some debate. Here, we present the results of a 12-y fieldworkproject, along with a taphonomic analysis of the human remains,designed to assess the funerary context of the La Chapelle-aux-Saints Neandertal. We have established the anthropogenic natureof the burial pit and underlined the taphonomic evidence ofa rapid burial of the body. These multiple lines of evidence supportthe hypothesis of an intentional burial. Finally, the discovery ofskeletal elements belonging to the original La Chapelle aux Saints1 individual, two additional young individuals, and a second adultin the bouffia Bonneval highlights a more complex site-formationhistory than previously proposed.

Mousterian burial | Middle Paleolithic | symbolic behavior | taphonomy |archaeology

Since the discovery of the La Chapelle aux Saints skeleton in1908 (LCS1) (refs. 1–4 and SI Appendix, Fig. S1), intentional

primary Neandertal burials have often been considered as keyevidence of behavioral modernity (5, 6). However, some scholarshave remained skeptical (7), arguing that most of these specialtreatments of the dead were identified in the context of old andinadequate excavations. During the past decade, possible evi-dence of Neandertal symbolic behavior has been claimed, in-cluding the use of pigment (8) and decorative feathers (9), thecollection of exogenous fossil shell (10), and a preference forcolored shells in Middle Paleolithic layers (11). In light of thisevidence, reassessment of the burial question is needed to bettercharacterize the Neandertal cultural package. Here, we reporton archaeological evidence from the La Chapelle-aux-Saintsburial, derived from taphonomic analysis of the LCS1 skeleton,as well as the study of the geological context of the deposit.These data provide insights relevant to the archaeological con-text of the Paleolithic human occupations at La Chapelle-aux-Saints, and they support previous claims for the existence ofintentional burial. By so doing, they buttress claims for complexsymbolic behavior among Western European Neandertals.

The Bouyssonie Discovery and Ancient Evidence forNeandertal BurialOn August 3, 1908, the Bouyssonie brothers discovered a nearlycomplete Neandertal skeleton (LCS1) in a pit dug within thedeposits of the bouffia Bonneval. For the first time, the hy-pothesis of the possible existence of intentional burials, and,therefore, symbolic capacities in an Upper Pleistocene humangroup other than anatomically modern humans, was clearly

raised (1). Consequently, our approach to understand prehistorichuman groups was drastically modified and archaeologists beganto look for Neandertal burial evidence. In the 5 y following thediscovery at La Chapelle-aux-Saints, 9 more purported burialswere discovered, and today nearly 40 possible cases have beenreported (12), some of which (Kebara 2 and Shanidar 4/6/8/9)reflect complex funeral practices (13, 14). All of these discoverieshave profoundly changed our perception of Neandertals.However, in the last few decades, numerous criticisms and

doubts have been raised regarding the reality of some of theseburials, including questions as to whether Neandertals actuallypossessed the cognitive capability to bury their dead (7, 15). Interms of the bouffia Bonneval discovery, the lack of informationregarding the Bouyssonie’s excavation procedures has been usedas support for reservations concerning the burial hypothesis (7).Because the site’s formation processes remained relatively un-known, the agent responsible for the preservation of the skeletalremains was unclear, and the exact nature of the burial pit wasstill not firmly established. Similar questions have recently beenproposed with regard to the Roc-de-Marsal 1 Neandertal childburial (15), where a reassessment of the sedimentation processand the archaeological context of the skeletal remains have castdoubts on its intentional nature.

Significance

For several decades, scholars have questioned the existence ofburial in Western Europe prior to the arrival of AnatomicallyModern Humans. Therefore, an approach combining a globalfield recovery and the reexamination of the previously dis-covered Neandertal remains has been undertaken in the siteof La Chapelle-aux-Saints (France), where the hypothesis of aNeandertal burial was raised for the first time. This project hasconcluded that the Neandertal of La Chapelle-aux-Saints wasdeposit in a pit dug by other members of its group and pro-tected by a rapid covering from any disturbance. These dis-coveries attest the existence of West European Neandertalburial and of the Neandertal cognitive capacity to produce it.

Author contributions: W.R., C.B., T.B., and B.M. designed research; W.R., C.B., I.C., P.B.,T.B., L.B., G.D., J.-P.F., C.T., D.T., A.T., and B.M. performed research; A.B., F.L.-C., and C.T.contributed new reagents/analytic tools; W.R., C.B., I.C., P.B., L.B., G.D., J.-P.F., F.L.-C., D.T.,A.T., and B.M. analyzed data; and W.R., C.B., I.C., P.B., D.T., and B.M. wrote the paper.

The authors declare no conflict of interest.

This article is a PNAS Direct Submission.1To whom correspondence should be addressed. E-mail: williamrendu@hotmail.fr.

This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10.1073/pnas.1316780110/-/DCSupplemental.

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Considering the controversy surrounding the issue of Neandertalburials and problems raised concerning La Chapelle-aux-Saints,we developed a twofold approach incorporating a field projectcombined with the taphonomic analysis of the LCS1 skeletalmaterial.

Reevaluation of La Chapelle-aux-SaintsExcavations carried out along the cliff and at the entrance ofthe bouffia Bonneval were designed to gather new informationfor establishing a more general understanding of the prehistoricoccupation. Seven additional shallow north facing eroded cavi-ties (SI Appendix, Fig. S2 and ref. 16), formed at the interface ofthe Upper Hettangian limestone of the cliff and the LowerHettangian marl, were explored. In terms of morphology, theseclosed cavities appear as small rock shelters rather than well-developed karstic conduits. Features related to evolved karstdrainage system (i.e., water circulation) are absent on the wallsor the roof of the cavities, and the multiple test pits producedonly limited traces of endokarstic sediments. Therefore, shelterformation appears mainly connected to (i) the opening of thelimestone/marl interface of the Jurassic cliff (17), (ii) the retreatof the cliff by regressive erosion, (iii) the erosion and reworkingof autochthonous weathered sediments at the lithological in-terface in the cliff, and (iv) the trapping of allochtonous sedi-ments transiting along the slope.Each of the rock shelters yielded Late Middle Paleolithic and

Upper Paleolithic materials. A Quina Mousterian industry wasidentified in bouffia Bonneval and bouffia 102, LevalloisianMousterian was identified in bouffia Bonneval and bouffia 118,and Mousterian of Acheulean Tradition (MTA) was identifiedonly in bouffia 118. Additionally, Châtelperronian artifacts wererecognized in bouffias 102, 129, and 137–140, whereas severalUpper Paleolithic artifacts (Aurignacian or Magdalenian ones)were found in bouffias 102 and 129 (SI Appendix, Fig. S2). Thearchaeological sequence for the Middle-to-Upper Paleolithictransition in southwestern France is therefore represented at LaChapelle-aux-Saints (18).Major taphonomic processes affected the integrity of the

stratigraphic and archaeological units. Periglacial features (es-pecially cryoturbation) were observed during excavation of all ofthe bouffias. Furthermore, mixed deposits from hyena dens andhuman occupations were also recognized at the Grotte du Noyer,as well as at bouffias 102, 131–133, and 137–140 (SI Appendix,Fig. S2). Finally, because the cliff limestone was eroded andconsequently retreated, parts of the cavities’ deposits wereprobably affected and remobilized along the slope.Only bouffia 118 produced preserved archaeological strata:

the lower level (C2) is clearly dominated by reindeer and is as-sociated with a Denticulate Levallois Mousterian, whereas theupper layer (Alpha) is rich in bovines and is linked to a MTAindustry (SI Appendix, Text S1 and Table S1). These two occu-pations have been attributed, respectively, to marine isotopic stage(MIS) 4 and MIS 3 based on biochronological data (SI Appendix,Text S1 and Fig. S3). Moreover, the taphonomic analysis dem-onstrates clear differences in the preservation of the faunalmaterial. Whereas the C2 remains are systematically stained byFe-Mn oxide deposits, those from layer Alpha, although free ofthese modifications, are heavily weathered (SI Appendix, Table S2).This information provides important contextual insights for theMousterian occupations at the bouffia Bonneval.Excavations in the bouffia Bonneval took place in 2011 and

2012. Multiple test pits were placed in a 23.5-m2 area of the cave,as well as on the slope at the cave’s entrance (Fig. 1). Thesediments were backfill of the Bouyssonies’ excavations bothinside and outside, except for small areas where in situ depositswere identified inside the cave.The marl substratum was reached 10 cm below the surface

layer in the cave, and 40 cm on the slope outside under the

backfill deposits from the older excavations. The excavatedsediments were rich in Middle Paleolithic artifacts similar tothose previously recovered by the Bouyssonies. The assemblageis composed of numerous Levallois flakes and Quina scrapersmade from quartz, as well as from both local and nonlocal flint.Bifacial shaping technology is also occasionally documented andsuggests the presence of at least two Mousterian components(Quina and MTA). In the entire archaeological record of south-western France, these two different chaînes opératoires are neverfound in association and appear not to be contemporaneous(18–20).The faunal spectrum is dominated by reindeer, followed by

bovine, with carnivores being represented by wolf, fox, andbadger (SI Appendix, Table S3). Reindeer and bovine remainsexhibit two different preservation patterns; natural oxide color-ation is significantly more frequent on the reindeer elementsthan on bovine material, which in turn exhibits a higher fre-quency of weathering modifications. These features stronglysuggest two different taphonomic histories for the two faunalcomponents. The combined results of the lithic and faunal anal-yses support the hypothesis of two distinct Mousterian levels, onedominated by reindeer and the second mostly composed of bo-vine. As documented in the bouffia 118, the reindeer episodewould be the earlier one.Therefore, the recovering of several preserved (i.e., unexca-

vated by previous archaeologists) areas is of prime interest. Ina sector of a square meter surface, located in the west part of theentrance under a limestone collapse, an in situ deposit wasidentified. Laying directly on the marl substratum, this depositcorresponds therefore to the first recorded occupation of the

Fig. 1. Bouffia Bonneval excavation map and burial pit position. Here,differences in the cavity topography and in the localization of the burial pitare linked to imprecision in the Bouyssonies’ drawing.

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cavity (stratigraphic unit C4). It is characterized by homogenousarchaeological material composed of Quina Mousterian and a boneassemblage dominated by reindeer remains. The recurrent pref-erential orientation of the elongated remains and the very lowfrequency of recent fracture on the bone splinters attest thatthis area was not disturbed by the early 20th century excavation.Moreover, in at least two distinct areas within the cavity, mate-rials were identified trapped in the first centimeters of the sub-stratum (stratigraphic unit C2sup).As a consequence, considering the local (in situ deposit and

the bouffia 118) and the regional biochronological contexts (18),a stratigraphy can be inferred from the backfill analysis. TheQuina Mousterian assemblage, associated with a reindeer ex-ploitation, would have been followed by at least one other layerrelated to MTA Mousterian in association with a bovine ex-ploitation. An attribution of the two theoretical assemblages toMIS 4 and MIS 3, respectively, can be proposed. The reindeer/Quina Mousterian occupation would therefore have been re-worked by cryogenic processes that affected the marls (e.g., in-jection features).In addition to the artifacts, a total of 13 human elements were

unearthed during the recent excavations. Four elements belongto the LCS1 skeleton; the root of an upper second premolar,a fragment of a scapula, the distal extremity of an ulna, anda complete pollical proximal phalanx (SI Appendix, Text S2 andFigs. S4 and S5). These remains complement the previous dis-covery in the 1920s of an isolated upper right third molar (SIAppendix, Text S2 and Fig. S4) in the bouffia Bonneval (21).Moreover, three additional individuals have been recognized,

based on a left upper second premolar, a distal root of a lowerpermanent molar, a right upper central deciduous incisor, a rightupper deciduous second molar, and a right lower deciduous firstmolar. These remains attest to the presence of two children andanother mature individual (SI Appendix, Text S2).All of these remains fall within the morphological variability of

the Neandertals.

The Burial PitExcavations within the bouffia Bonneval exposed a 39-cm-deepsubrectangular depression cut into the substrate marl. The upperpart of the pit is damaged; the lower part is 140 cm long and 85cm wide (SI Appendix, Figs. S6–S8). The interface of the backfilland the substratum is clear and regularly concave in this area. Itsposition is identical to the location of the burial pit indicated inthe Bouyssonies’ figures (Figs. 2 and 3). This is also supported bythe relative position of the pit in relation to the relief of the roof(SI Appendix, Fig. S7) observable in the excavation pictures fromthe early 20th century.The lack of evolved karstic morphologies and sedimentation

connected to groundwater action in the bouffia Bonneval and inthe adjacent La Chapelle-aux-Saints loci reject the hypothesis (7)of an endokarstic origin for the depression. Furthermore, the pitdoes not have the same morphology as the cryoturbation cells,ruling out a periglacial origin. The pit is also unlikely to derivefrom a brown or cave bear hibernation nest. It does not have thesame morphology of such nests (refs. 22 and 23 and Fig. 2), ursidremains are quasi absent from the faunal spectrum (SI Appendix,Table S4) and its origin (natural or carnivore accumulation) islargely unclear, and the shallow depth and relatively largeopening of the bouffia Bonneval make it unlikely to have servedfor bear hibernation (Figs. 1 and 3).It is not clear whether the hollow into the marl was excavated

by the resident Neandertals, a natural feature or a natural de-pression enlarged by them. No tool marks into the marl weredetected. However, the pit cut partially through a cryoturbatedfissure (stratigraphic unit C5) that contained Quina artifacts (Fig.4). The last aspect demonstrates that the area of the depressionwas at least partially modified for the burial of LCS1, removing

accumulated cultural debris and sediment. This modificationwould have happened after the formation of the cryoturbatedfissure, thus after the Quina deposit. Therefore, based on theexcavations along the La Chapelle-aux-Saints cliff, an anthro-pogenic origin for the depression that contained the articulatedremains of LCS1 at the bouffia Bonneval is the most parsimo-nious explanation.

Taphonomic Patterns of the LCS1 NeandertalThe taphonomic analysis of the human [number of identifiedspecimens (NISP), 77] and faunal (NISP, 1,533) remains re-covered in the early 20th century has included the systematicrecording of climatic, mechanical, and carnivore modificationsvisible on the bone surfaces. The preservation of the humanbones is significantly different from the pattern observed for thefaunal material. The cortical surfaces of the Neandertal bonesare significantly less deteriorated than the reindeer (χ2: 307.3;P < 0.001) or bovine (χ2: 212.7; P < 0.001) material. Only longi-tudinal cracks are observable on the LCS1 remains and smoothingand exfoliation are absent, whereas such modifications are fre-quent on the faunal elements (SI Appendix, Tables S5 and S6).No carnivore marks are observable on the human bones, al-though their occurrence on the faunal material (carnivore marks:4.2% NISP) indicates that carnivores had access to the discardedcarcasses.These data highlight two different taphonomic histories for

the human and faunal material, lending considerable support tothe hypothesis that the Neandertal corpse was covered rapidly.Furthermore, all major anatomical regions are represented by

Fig. 2. Pit profile. The letters refer to Fig. 1. The in situ remains, found inthe cryoturbated cell and observable in the A–B profile are in yellow. Allother remains were found in the backfill.

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the human skeletal remains (SI Appendix, Fig. S9), despite theabsence of many of the smaller skeletal elements.The recurrent modern fractures on all of the long bones of the

LCS1 reflect the speed at which the Bouyssonies conducted theirexcavations and not preexcavation processes.

DiscussionRecent excavations at La Chapelle-aux-Saints have revealedMiddle Paleolithic and carnivore occupations in numerous cav-ities along the cliff (17). Thousands of lithic artifacts and bonefragments attest to long-term occupations during MIS 4-3. Thetaphonomic recovery and our excavations in the bouffia Bon-neval argue in favor of the existence of at least two distinct ar-chaeological assemblages. After the deposition of the QuinaMousterian layer during the MIS 4, some cryoturbation oc-curred, injecting part of the material within the substratum. Atsome stage in the MIS 3, a group of bovine hunters, related tothe MTA complex, occupied the cavity. Throughout these, atleast two, occupations, Neandertals introduced a large numberof medium and large ungulates, namely bovine and reindeer,which were processed on-site. Moreover, the excavations reportedhere have identified the exact position of the LCS1 burial pit.The results of the comparative taphonomic analysis of the hu-

man and faunal materials demonstrate that the LCS1 Neandertal

corpse was rapidly interred and protected from the post-depositional modifications experienced by the faunal remains.The existence of an artificially modified pit and the rapidburial of the body constitute convincing criteria for estab-lishing purposeful burial during the Middle Paleolithic ofWestern Europe.Whether the origin of the pit is natural or anthropogenic is not

considered here as a discriminatory factor because the oppor-tunistic use of natural depressions in funeral practices has al-ready been demonstrated in Upper Paleolithic contexts (24) andtherefore cannot be ruled out as a possibility. The lack of in-formation about the original stratigraphy and excavations makesit impossible to address the dynamic sedimentary processes in-volved in the filling of a pit. However, three arguments supportthe rapid burial of the corpse: the completeness of the cranialand infracranial elements, the existence of anatomical connections,and the preservation patterns seen on the cortical surfaces ofthe bones.In Western Europe, 20 primary or secondary burials (25) have

been proposed (SI Appendix, Table S7) for Neandertals. In eachcase, the burial hypothesis is based on at least one of these twocriteria: the presence of a pit and the rapid burial of the remains.However, only 8 cases exhibit the two criteria in conjunction, andfor the 12 other potential burials, the available information isinsufficient for the burial context to be properly evaluated.It should be highlighted that in every case where sufficient

data are available, the burial is associated with an occupationcharacteristic of a residential camp (26, 27). Therefore, no task-specific location (28) in the European Middle Paleolithic can belinked exclusively to funerary activities (29). It thus seems thatsymbolic manifestations and economic patterns are both firmlyembedded within European Neandertal territories.

Fig. 3. The bouffia Bonneval in 1909 and September 2012. The 1909 picturecomes from the Fonds Bouyssonie-Ecole Bossuet.

Fig. 4. Cryoclastic figure cut by the pit (stratigraphic unit C5).

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ConclusionMore than a century after the discovery of the La Chapelle-aux-Saints skeleton, we have corroborated the information providedin the original excavation reports concerning the finding of anarticulated, complete human skeleton within a depression in thebedrock. Microstratigraphic observation of the edges of the de-pression indicates that it postdates both the accumulation ofQuina Mousterian deposits and their postdepositional cryoturba-tion and, therefore, that, originally, it cut through sedimentfill, first, and then the bedrock itself. The anthropic origin of theexcavation of this feature is the parsimonious reading of the evi-dence; a geogenic origin can be excluded, and there is no evidencethat cave bears used the site for hibernation (and the site is tooshallow for that to be possible in the first place). The taphonomyof the human remains sets them clearly apart from the site’s fauna,because no carnivore modification is apparent, indicating rapidburial, as one would expect in a funerary context. No reason existsto question the interpretation of the LCS1 burial.The results include the identification of three further Nean-

dertal individuals, additional elements attributed to the first in-dividual, as well as several other shelters occupied by Middle andUpper Paleolithic human groups.In the light of our work at La Chapelle-aux-Saints and fol-

lowing the recent revision of the Roc-de-Marsal child burial, itnow appears that a general reassessment of European Mousterianburials needs to be undertaken with the aim of furnishing newscientific arguments and evidence relevant to the ongoing debatesurrounding Neandertal symbolic behavior.Whatever the status may be of other purported Neandertal

burials, the evidence from the bouffia Bonneval at La Chapelle-aux-Saints serves to establish the existence of intentional Neandertalinhumation in the European Middle Paleolithic. Therefore, thiswork contributes to the current redefinition of the general pictureof Neandertal cultural behavior.

Materials and MethodsExcavation Method. From 1999 to 2010, we excavated seven cavities at LaChapelle-aux-Saints. One of them, bouffia 118, provided a significant series oflithic and animal remains (17). From 2011 to 2012, the excavations werecentered in the Bouffia Bonneval. During 2011 excavations, only backfilldeposits were identified in the cavities. In 2012, several in situ deposit wereidentified and excavated. At the end of the field project, more than 80% ofthe surface of the cavity was excavated. The excavations followed the sameprotocol during all of the process, whatever the nature of the excavatedsediment (backfill or in situ deposit). All lithic artifacts larger than 25 mmand fauna larger than 30 mm were given 3D coordinates with a total station.Identifiable bones and every macrofaunal tooth fragment smaller than 30mm were systematically collected. All natural stones larger than 10 mm wererecorded with a single coordinate, and all natural stones larger than 150 mmwere measured with multiple coordinates to describe their volume. A firstanatomical and taxonomical identification was undertaken on the field foreach faunal remain to identify in situ the potential human remains.

Sediment was collected by stratigraphic units and by quarter of squaremeter surfaces every 5-cm depth and was water-screened through finemesh (1.8 and 2.2 mm). The coarse fraction was sorted to separate lithics,fauna, and any other archaeological items. Human remains were methodi-cally looked for in the coarse fraction in a laboratory by a human pale-ontologist expert. Every cavity, locus and stratigraphic unit is described indetail in the excavation report that was presented to the Department ofArchaeology of the Limousin Region (Service Régional de l’Archéologiedu Limousin) on February 8, 2012*.

Taphonomic Analysis. The analysis concerned the faunal remains excavated inbouffia 118 and in bouffia Bonneval during the recent field project (morethan 8,500 remains), the faunal material excavated during the early 20thcentury [Number of Remains (NR), 1,841], and the totality of the Neandertalremains discovered by the Bouyssonies (NR, 77) and located in the Musée del’Homme in Paris (France). Every piece from the different faunal assemblageswas analyzed by physical paleoanthropologist to recover all possible humanremains. Pieces were identified at the most precise level, and, when it wasnot possible to propose a specific attribution, ungulate size classes wereused (following ref. 30 modified from ref. 31). With regard to the skeletalpart profiles, all identifiable specimens (including shaft fragments) collectedwere taken into account and recorded following the “element, portion,segment” coding format established by Gifford and Crader (32). Analysisof the bone surfaces for the faunal and human remains excavated bythe Bouyssonies was performed by the same analyst (C.B.) to avoid in-terobserver variability. Weathering, root etching, anthropogenic, andcarnivore modifications were systematically looked for using the criterialisted by Blumenschine et al. (33) and others (34–37). Moreover, oxide col-orations of the bone cortical surfaces were carefully recorded. The stage ofpreservation of the cortical surfaces was estimated (following refs. 38 and39). When unclear modifications were detected, specimens were subjectedto more thorough evaluation with a 10–40× microscope. Percentagevalues were calculated on the basis of the total number of remains, boneswith unobservable surfaces having been excluded (following ref. 40).

ACKNOWLEDGMENTS. We thank Joao Zilhao, Brad Gravina, Lars Anderson,and Randall White for helpful comments. We thank the two anonymousreviewers and E.T. for valuable advice. We thank Mrs. Bouyt, Decoux, andLevet, owners of the site; Martine Fabioux (head of the Limousin archaeo-logical service); Aurélie Fort and Philippe Mennecier from Musée de l’Homme;and the Accès Scientifique à la Tomographie à Rayons X platform, Unité Mixtede Service 2700 (Outils et méthodes de la systématique intégrative), CentreNational de la Recherche Scientifique–Muséum National d’Histoire Naturelle(Paris) for the micro-CT scanning procedures. This work has been supportedby the French Ministry of Culture; the Archéosphère Company; Unité Mixtede Recherche Travaux et Recherches Archéologiques sur les Cultures, lesEspaces et les Sociétés (UMR TRACES) (Toulouse, France); Préhistoire à l’ActuelCulture, Environnement et Anthropologie (Bordeaux, France); the La Chapelle-aux-Saints municipality and its inhabitants; and the Communauté de Com-munes du Sud Correzien. We thank the SMP3C team of the UMR TRACESfor translation funding. W.R. was funded by the Fyssen Foundation andsupported by the University of Michigan.

*Rendu W, Beauval C, The Bouffias Excavation at La Chapelle-aux-Saints, Report to theArchaeological Regional Service of Limousin of Antiquities, February 8, 2012, Limoges, France.

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22. Fosse P, Besson JP, Laborde H, Thomas-Cantie F, Cazenave G, Delmasure MC,Leveque T, Laudet F, Quiles J (2004) Denning behaviour of “modern” brown bear(Ursus arctos, L.) in caves: Biological and paleontological considerations from FrenchPyrenean sites. Cah Sci Mus Hist Nat Lyon 2:171–182.

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25. Duday H (2011) The Archaeology of the Dead, Lectures in Archaeothanatology(Oxbow Books, Oxford).

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28. Binford L (1980) Willow smoke and dogs’ tails: Hunter-gatherer settlement systems

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from Soleihac (Haute-Loire) and from Lenel-Viel 1 (Herault)] (Université de Provence,

Aix-Marseille I, Aix-en provence, France). French.31. Brain CK (1981) The Hunters and the Hunted? An Introduction of African Cave Ta-

phonomy (Univ of Chicago Press, Chicago).32. Gifford DP, Crader DC (1977) A computer coding system for archaeological faunal

remains. Am Antiq 42:225–238.33. Blumenschine RJ, Marean CW, Capaldo SD (1996) Blind tests of inter-analyst corre-

spondence and accuracy in the identification of cut marks, percussion marks, and

carnivore tooth marks on bone surfaces. J Archaeol Sci 23:493–507.34. Olsen SL, Shipman P (1988) Surface modification on bone: Trampling versus butchery.

J Archaeol Sci 15:535–553.35. Pickering TR, Egeland CP (2006) Experimental patterns of hammerstone percussion

damage on bones: Implications for inferences of carcass processing by humans.

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ering. Paleobiology 4:150–162.37. D’Errico F, Villa P (1997) Holes and grooves: The contribution of microscopy and ta-

phonomy to the problem of art origins. J Hum Evol 33(1):1–31.38. Rendu W (2007) Planification des activités de subsistance au sein du territoire des

derniers Moustériens. Cémentochronologie et approche archéozoologique de gise-

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supérieur ancien (Isturitz) [Subsistence Activity Scheduling Within the Territory of the

Last Mousterians Skeletochronological and Zooarchaeological Approaches Applied to

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laeolithic One (Isturitz)] (Université Bordeaux 1, Talence, France). French.39. Rendu W (2010) Hunting behavior and Neandertal adaptability in the Late Pleisto-

cene site of Pech-de-l’Azé I. J Archaeol Sci 37:1798–1810.40. Delpech F, Villa P (1993) Exploitation des Animaux Sauvages á travers le Temps [Wild

Fauna Exploitation Through Time], eds Desse J, Audoin-Rouzeau F (Association pour

la Promotion et la Diffusion des Connaissances Archéologiques–Centre National de la

Recherche Scientifique, Sophia Antipolis, France), pp 79–102.

6 of 6 | www.pnas.org/cgi/doi/10.1073/pnas.1316780110 Rendu et al.

One hundred years later: new evidence supporting an intentional Neandertal

burial at La Chapelle-aux-Saints (Corrèze, France)

William Rendu, Cédric Beauval, Isabelle Crevecoeur, Priscilla Bayle, Antoine Balzeau,

Thierry Bismuth, Laurence Bourguignon, Géraldine Delfour, Jean-Philippe Faivre, François

Lacrampe-Cuyaubère, Carlotta Tavormina, Dominique Todisco, Alain Turq, Bruno Maureille.

SUPPORTING INFORMATION (SI) SUPPORTING TEXTS ............................................................................................................................. 2 

TEXT S1: Bouffia 118 zooarchaeological analysis ............................................................................ 2 

TEXT S2: New Neandertal remains discovered during the 2011‐2012 excavations ....................... 2 

SUPPORTING FIGURES ......................................................................................................................... 7 

FIGURE S1: Site location and picture of the site entrance. ............................................................. 7 

FIGURE S2: Map of the La Chapelle‐aux‐Saints cavity complex and main stratigraphic 

information. ..................................................................................................................................... 8 

FIGURE S3: Mousterian sites with faunal spectra dominated by Bison or Reindeer attributed to 

the MIS 4 or the MIS 3 ..................................................................................................................... 9 

FIGURE S4: Human dental remains discovered during the 2011‐2012 excavations and during the 

1920’s. ........................................................................................................................................... 10 

FIGURE S5: Human osseous remains discovered during the 2011‐2012. ..................................... 11 

FIGURE S6: First excavation stage of the pit structure in 2011. .................................................... 12 

FIGURE S7: Picture of the excavators nearby the burial pit, after its excavation ......................... 12 

FIGURE S8: Picture of the pit after the 2012 excavation. ............................................................. 13 

FIGURE S9: Skeletal part representation of LCS1 after the 2011‐2012 discoveries. ..................... 14 

SUPPORTING TABLES ......................................................................................................................... 15 

TABLE S1: Faunal spectrums from Bouffia 118. ............................................................................ 15 

TABLE S2: Taphonomic modifications of the Alpha and C2 faunal remains from Bouffia 118. .... 15 

TABLE S3: Faunal spectra of the Bouffia Bonneval from the 2011‐2012 excavations. ................. 16 

TABLE S4: Faunal spectrum of the Bouffia Bonneval from the Bouyssonies’ excavations. .......... 17 

TABLE S5: Taphonomic alterations of the faunal remains from the Bouyssonie excavation. ...... 18 

TABLE S6: Taphonomic alterations of the faunal remains from the recent excavation. .............. 18 

TABLE S7: Potential primary and secondary Western European Neandertal burial. .................... 19 

REFERENCES FOR SUPPORTING INFORMATION ................................................................................ 20 

SUPPORTINGTEXTS

TEXTS1:Bouffia118zooarchaeologicalanalysis

Two archaeological levels have been identified in Bouffia 118: Alpha and underlying C2.

Alpha has yielded a large quantity of bifacial thinning flakes linked to the MTA techno-

complex (see 1). The faunal assemblage from level Alpha is largely dominated by Bison

remains (Table S1) presenting a high frequency of weathering modifications and root

etchings (Table S2). All the skeletal elements seem to have been introduced into the site. On

the other hand, the faunal spectrum of layer C2 is largely specialized on reindeer (Table S1)

and is associated with a Levallois technology produced on local and exogenous raw

materials. While deer remains are very well-preserved, they are systematically stained by

natural manganese deposits (Table S1). Skeletal part profiles suggest a highly selective

pattern with more than 90% of the NISP being limb fragments. A statistically significant

relationship exists between the skeletal part representation and the quantity of marrow and

grease contained within the different imported elements.

Although the material is yet to be radiocarbon dated, comparisons with other well-dated

sequences from Southwestern France containing the same faunal associations allowed a

biochronological approach to the two layers (Fig S3). During the Middle Paleolithic of

Southwestern France, reindeer dominated faunal spectrums occur only during MIS 4 and the

very beginning of MIS 3 and are followed by a 'Bison phase' at around 45ky represented by

an overwhelming dominance of this large bovine in faunal assemblages recovered from sites

attributed to this period (for a complete synthesis see 2).

TEXTS2:NewNeandertalremainsdiscoveredduringthe2011‐2012excavations

Thirteen new hominin remains have been discovered during our field work in 2011 and 2012.

Seven of them are attributed to the LCS 1 individual based on their morphology and the

preservation of the skeleton. Additional dental remains, an upper third molar from the

collections of the Musée de l’Homme (Paris, France), a molar root fragment, and an upper

second premolar, could belong to additional adult individuals (18). Finally, three deciduous

teeth represent additional juvenile specimens.

New LCS 1 remains:

1) A right upper second premolar (LCAS 2011 R73 # 12) - Fig S4A&B

This tooth is limited to the root (S4). The virtual extraction of the left upper first premolar on

LCS 1 shows that this tooth is two rooted and exhibits a taurodont morphology. The

morphological similarities between the tooth still in place on the LCS 1 maxilla and LCAS

2011 R73 # 12 doesn't exclude the association of the new dental root to the same individual.

Fig S4B provides a virtual extraction and reconstruction of the first premolar still in place on

the left upper dental arch of LCS 1 allowing a comparison with LCAS 2011 R73 # 12.

2) A fragment of a right scapula (LCAS 2011 S73 #322) - Fig S5A

The piece is part of a right mature scapula preserving most of the glenoid cavity, 3

centimeters of the axillary border and 70 mm2 of the adjacent body. The roots of the coracoid

and acromion processes are broken. The morphology of the broken parts indicates ancient

breaks. As scapular elements are missing from the LCS 1 individual, we consider that the

simplest hypothesis is that this scapula fragment belongs to the LCS 1 skeleton. Note the

clear insertion of the M. teres minor on the dorsal surface (Fig S5A).

3) A distal extremity of a right ulna (LCAS 2011 S73 #62) - Fig S5B

This piece is 33 mm long and represents a well- preserved radial articulation with part of the

diaphysis. The proximal part of the fragment has an irregular and recent break. The antero-

posterior and transverse diameters of the articular extremity are respectively of 23.7 mm and

22.1 mm. The minimal circumference of the diaphysis, measured just above the radial

articulation, is 33.5 mm and the maximum height of the articular circumference is 9.5 mm.

No direct refitting is possible between the broken distal extremity of the right LCS1 ulna and

the newly discovered fragment as the original right ulna was repaired using different kinds of

materials. Nevertheless, the shapes of the two pieces suggest that this distal extremity

derives from the same bone.

4) A distal extremity of the left third metacarpal (LCAS 2012 S73 #449) - Fig S5C

This piece is part of the distal extremity of a metacarpal. The radial part of the articular

surface is well-preserved, while the palmo-ulnar side is broken. The comparison of this distal

extremity with the original material allows to attribute this fragment to the left third metacarpal

of LCS1, whose distal extremity is missing and reconstructed. The size and morphology of

the new fragment correspond with this missing area. Moreover, there is a perfect secondary

anatomical connection with the proximal articular surface of the right third proximal phalanx.

Finally, a shallow ovoid depression is present on the articular surface. The same

characteristic can be observed on the distal articular surface of the right second metacarpal

from the original collection.

5) A proximal phalanx of the right thumb (LCAS 2011 R74 #109) - Fig S5D

This piece is 30.7 mm long and is well-preserved with the exception of the palmar surface of

the diaphysis which is broken. Two small fragments of compact bone from this area are

however still preserved. The phalanx is slender and the lateral crests are not marked which

is typical for most of the Neandertal proximal pollical phalanxes (19). No pathological

conditions are perceptible.

Following the morphology and size of the other hand phalanxes associated to LCS 1

individual (20), we consider that the simplest hypothesis is that this piece belongs to the

same specimen.

6) A distal phalanx of the right thumb (LCAS 2012 S73 S2 Rem 25-30 seau 2) – Fig S5E

This piece is partially broken with the medial side of the apical tuft and part of the diaphysis

missing. Given the taphonomy of the break, it could results from the 1908's excavation. This

phalanx presents a large proximal extremity in relation to its maximum length following the

Neandertal pattern (19). The insertion area of the muscle pollicis longus is well marked. Only

the apical tuft shows few Neandertal affinity with a gracile distal extremity. Finally, the

proximal articular surface fits perfectly with distal articulation of the new right proximal

phalanx.

7) A distal extremity of the left third metatarsal (LCAS 2012 S73 #568) – Fig S5F

This piece is highly deformed by bone remodelling. It is however possible to identify it as the

distal extremity of a metatarsal. The articular surface is eburnated and flattened. Its

comparison with the original collection allowed to bring it closer to the right third metatarsal.

The symmetry is not straightforward because of the pathological state of this distal extremity,

but its dimension and morphology closely resemble the antimeric bone from the original

collection.

Presumed new adult specimens:

1) A left upper second premolar (LCAS 2012 R73 #221) - Fig S4C

This tooth presents a mesio-distal fracture and its root is apically broken. The crown shows a

very high wear degree, the lingual aspect of the enamel being completely worn and the pulp

cavity open. Only the maximal bucco-lingual diameter of the root can be estimated to 11 mm.

Compared to the right upper second premolar LCAS 2011 R73 # 12 likely belonging to

LCS1(see above), this new tooth has a shorter root with a morphologically different apex.

Thus, this tooth does not belong to LCS1, and represents a new adult individual at La

Chapelle-aux-Saints.

2) A right upper third permanent molar discovered in 1920 – Fig S4D

This right upper third molar was discovered September 7th, 1920 by a cousin of Bouyssonie

visiting the bouffia Bonneval who entrusted it to Jean Bouyssonie. It was recognized by the

latter as a human tooth. As he suspected it might belong to the LCS 1 fossil, he sent it to H.

Breuil (letter available at the Central library of the Muséum national d’Histoire naturelle, Paris

- Henri Breuil archives - classification mark Br 28) to give it to M. Boule. The tooth has been

in the fossil collections of the Musée de l’Homme (Paris) ever since (18).

Unfortunately, the upper alveolar arch of LCS 1 is poorly preserved and it is not possible to

test if this large third molar could refit.

3) A distal root of a lower permanent molar (LCAS 2011 R73 #26) - Fig S4E

This root of a human permanent molar bears a recent fracture on one face. The outward

orientation and distal curvature indicate that this root is a distal one of a lower molar. The

asymmetry of the lingual and the buccal edges (the buccal one is higher and less convex

than the lingual one) and the orientation of the apex suggest a left root. The slightly oblique

angle of the root relative to the inferred occlusal or cervical plane suggests that this root

cannot be a third molar and is probably more likely to be a second or a first molar.

This root belongs to either an adult or a child older than 12-13 years. There is substantial

resorption of the back part of the alveolar bone on both sides of the LCS 1 mandible. Such a

resorption is associated with a severe parodontopathy. No lower permanent molar would still

have been in place in the mouth of this Neandertal for at least several months before its

death. Thus, this fragment of a root of a lower left permanent molar cannot be attributed to

LCS1 and therefore represents another individual, like the one represented by the left upper

second premolar LCAS 2012 R73 # 221 (see above).

New juvenile specimens:

1) A right upper deciduous central incisor (LCAS 2011 S73 Rem 10-15 seau 1) – S4F

This tooth presents a complete functional crown and an apically broken root. The labial

surface is convex in both the vertical and horizontal planes. The mesial border is less convex

than the distal one which is also shorter. The lingual surface is shovel-shaped presenting a

well-developed mesial (higher) and distal (wider) margin and a clear lingual tubercule. The

mesial margin almost reaches the cingulum while the distal one is separated from it by a

deep surface. The crown occlusal border presents light attrition and there is also a very small

mesial interproximal contact facet at the mid-height of the crown. Considering the occlusal

wear, this tooth could have belonged to a child, circa 3 years old, younger than the Roc-de-

Marsal 1 and older than the Pech-de-l’Azé 1 specimens (21, 22). The crown mesio-distal

diameter is 8.17 mm and the bucco-lingual is 6.25 mm.

2) A right upper deciduous second molar (LCAS 2012 T74 S3 Rem 15-20 seau 1 & T73

S2 Rem 10-15 seau 2) - Fig S4G

This new tooth consists of two fragments from adjacent squares, one from T74 representing

the three quarters of the lingual crown, and the other one a small buccal portion from T73.

There are large fractures between the two fragments. The crown shows high occlusal wear

(6-1 following 23), and large interproximal contact facets. The beveled edge of the small root

fragment suggests that the tooth has been lost during the life of the individual. Following 24,

this tooth could have belonged to a 11-12 years old child. The crown mesio-distal diameter is

estimated to 8.83 mm, and the bucco-lingual to 11.50 mm.

Given their different wear patterns, this new tooth and the right upper deciduous central

incisor LCAS 2011 S73 Rem 10-15 seau 1 (see above) cannot have belonged to the same

juvenile individual.

3) A right lower deciduous first molar (LCAS 2012 Q74 S1 Rem 50-55 seau 1) - Fig S4H

This is a well-preserved crown showing a high occlusal wear (5-2 following 23), and

pronounced interproximal contact facets. A small fossa on the mesial aspect of the

protoconid recalls the morphology shown by Rescoundudou 2 (25), and Pech-de-l'Azé 1

(26). Following 24, this tooth could have belonged to a 10 years old child. The crown mesio-

distal diameter is 8.18 mm, and the bucco-lingual is 7.67 mm.

Given its wear degree, this new tooth could belong to the same individual as the right upper

deciduous second molar LCAS 2012 T74 S3 Rem 15-20 seau 1 & T73 S2 Rem 10-15 seau

2.

At least two juvenile individuals are recognized at La Chapelle-aux-Saints, one represented

by an upper deciduous central incisor, and the other one by an upper deciduous second

molar and a lower deciduous first molar.

SUPPORTINGFIGURES

FIGURES1:Sitelocationandpictureofthesiteentrance.A: Map From Geoatlas. B: the picture was taken from the base of the slope in front of the cave entrance before the 2011 excavation.

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FIGURES3:MousteriansiteswithfaunalspectradominatedbyBisonorReindeerattributedtotheMIS4ortheMIS31: Saint-Césaire, Layer EGPF (3); 2: Marillac-Les Pradelles, Layers 9 and 10 (4); 3: Jonzac, Layers 22, 20, 18, 16, 14, 12 and 9 (5-6); 4: La Quina, Layer 8 (7), Layers 6a and 6c (8-9); 5: Roc-de-Marsal (10); 6: Combe Grenal, Layers 17 and 16 (11); 7 : Pech de l’Azé IV, Layers I and H (12); 8: Sous-les-Vignes (13); 9: Puycelsi-La Rouquette, Layer 1 (14); 10: Mauran, (15-17). Cultural and radiochronological attribution are discussed and synthesized by (2).

FIGURES4:Humandentalremainsdiscoveredduringthe2011‐2012excavationsandduringthe1920’s.A, C, E-H dental remains discovered during the 2011-2012 excavation. D: M3 discovered in 1920. B : virtual extraction and reconstruction of the first premolar still in place on the left upper dental arch of LCS 1 allowing a comparison with LCAS 2011 R73 # 12

FIGURES5:Humanosseousremainsdiscoveredduringthe2011‐2012.

FIGURES6:Firstexcavationstageofthepitstructurein2011.The dark sediment corresponds to the Bouyssonie’s backfill; the light corresponds to the marl. The dotted line marks the pit limit.

FIGURES7:Pictureoftheexcavatorsnearbytheburialpit,afteritsexcavationLeft: Jean Bouyssonie and Father Robert visiting the site just after the discovery; Right: current excavators in 2011. The white line on the two pictures underlines the same relief of the cavity roof. The pictures were taken from the entrance of the cavity. The 1909 picture comes from the Fonds Bouyssonie-Ecole Bossuet.

FIGURES8:Pictureofthepitafterthe2012excavation.Backfill can only be seen now at the extremity of the cavity and the substratum is exposed on the totality of the excavated area. The picture was taken from the entrance of the cavity in September 2012.

FIGURES9:SkeletalpartrepresentationofLCS1afterthe2011‐2012discoveries.Grey: Skeletal elements identified in 1908; Red: Skeletal elements from the 2011 excavation. Drawing from: URA 376 CNRS after S.T. Constandse - Westermann and C. Meikeljohn, modified by M. Guillon, P. Sellier and P. Courtaud. Digitized by M. Coutureau (INRAP).

SUPPORTINGTABLES

TAXA ALPHA C2

Fox 7 2

Hyena 1

Wild cat 1

Reindeer 35 605

Roe deer 1

Cervid 1

Bovine 140 78

Horse 2

Avifauna 3

Marmotte 1 1

NISP 189 689

Medium sized ungulates 2 11

Large sized ungulates 226 73

Mammals 50 46

Total 467 819

TABLES1:FaunalspectrumsfromBouffia118.Faunal spectra established on the two main layers from bouffia 118. Alpha is the upper layer, dominated by bovine remains; C2 is the lower layer, dominated by reindeer.

Taphonomic

modifications

Alpha C2

NR %NR NR %NR

Longitudinal cracks 104 22,27% 237 28,94%

Exfoliation 163 34,90% 169 20,63%

Flaking of the outer

surface 65 13,92% 72 8,79%

Smoothed bone 98 20,99% 58 7,04%

Manganese deposits 11 2,36% 115 14,05%

Carnivore modifications 26 5.07% 51 6.23%

TABLES2:TaphonomicmodificationsoftheAlphaandC2faunalremainsfromBouffia118.Significant differences can be observed in the preservation pattern between the upper layer (Alpha) and the lower one C2 particularly if we considered the frequencies of smoothed bones and manganese deposits.

Taxa Backfill C4 C2sup C5 Wolf 3 1 Fox 61

Hyena 3 Felid 3

Badger 60 Polecat 0 1

Carnivore 2 Hyduntinus 0 1

Horse 56 7 3 Ovicapra 9

Capra 1 Bovine 675 87 1 1

Roe deer 5 3 Reindeer 4225 439 98 19 Red deer 7 1 Giant deer 1 Wild boar 13 Marmot 8 7

Lagomorph 101 NISP 5233 538 111 20

Large size ungulate 751 84 3 4

Mammal 457 36 2

Avifauna 78 NR 6519 658 114 26

TABLES3:FaunalspectraoftheBouffiaBonnevalfromthe2011‐2012excavations.C4: material from the non previously excavated area; C2sup: material moved into the marl by the cyroturbation. C5: faunal remains from the cryoclastic feature cut by the pit. The Backfill assemblage corresponds to what had been discarded by the Bouyssonies during their excavation.

Bouyssonies' excavation TAXA NISP Bear 1 Wolf 1 Fox 4

Horse 25 Rhinoceros 3 Ovicapra 5

Capra 3 Bovine 915

Reindeer 529 Red deer 12 Giant deer 8

Cervid 19 Wild boar 1 Marmot 1

Lagomorph 6 NISP 1533

Large size ungulate 255

Mammal 53 NR 1841

TABLES4:FaunalspectrumoftheBouffiaBonnevalfromtheBouyssonies’excavations.We have here to keep in mind that the sampling was highly selective and large taxa might be over-represented in this collection. It probably explains the main differences between the Bouyssonies collection and the material yielded by the backfill excavation.

Taphonomic alterations Reindeer Bison Neandertal NR % NR % NR %

Longitudinal craks 220 41,59 439 47,98 5 6,49 exfoliation 233 44,04 241 26,33 0 0 freezing modification 46 8,69 116 12,68 0 0 smouthed bone 196 37,05 180 19,68 4 5 Manganeze deposit 42 7,94 34 3,72 6 8

Bonesurfacealteration0 - 1 260 55,32 85 41,67 77 100

Bonesurfacealteration2 89 18,94 60 29,41 0 0

Bonesurfacealteration3 75 15,96 45 22,06 0 0

Bonesurfacealteration4 46 9,79 14 6,86 0 0

Carnivoremodifications 21 4 47 5,1 0 0

TABLES5:TaphonomicalterationsofthefaunalremainsfromtheBouyssonieexcavation.Bone surface alteration pattern refers to the stage of preservation of the cortical surface of the remains (following (16)). 0-1: less than 25% of the remain surfaces are altered, 2: less than 50% of the remain surfaces are altered. 3: less than 75% of the remain surfaces are altered, 4: more than 50% of the remain surfaces are altered. Major differences can be observed between the different analyzed taxa. Human remains are significantly less altered than the faunal ones. Bison and reindeer exhibit also two different patterns of preservation: with for instance frequencies of exfoliation and manganese deposits higher on reindeer remains than on bison ones.

Taphonomic alterations Reindeer Bison NR % NR %

Longitudinal craks 150 11,82 46 20,91 exfoliation 418 32,94 73 33,18 freezing modification 111 8,75 17 7,73 smouthed bone 81 6,36 18 8,18 Manganeze deposit 180 32,07 19 19,09 Bone surface alteration 0 - 1 730 62,07 115 59,89 Bone surface alteration 2 211 17,94 32 16,67 Bone surface alteration 3 99 8,42 23 11,98 Bone surface alteration 4 136 11,56 22 11,46 Carnivore modifications 30 2,4 9 4,1

TABLES6:Taphonomicalterationsofthefaunalremainsfromtherecentexcavation. Sample made from the 2011 material. Differences between the two collections might be explained by the fact that part of the recent one is composed by very small fragments which have a smaller cortical surface and, thus, have a lower probability to exhibit carnivore marks.

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